BAKTERI ANAEROB

MORFOLOGI, FISIOLOGI,
EPIDEMIOLOGI, DIAGNOSIS,
PEMERIKSAAN
Sy. Miftahul El J.T

Figure 4-1 Catabolism of proteins, polysaccharides, and lipids produces glucose, pyruvate, or
intermediates of the tricarboxylic acid (TCA) cycle and, ultimately, energy in the form of adenosine
triphosphate (ATP) or the reduced form of nicotinamide-adenine dinucleotide (NADH).
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Oksigen berdasarkan keperluan oksigen dibagi dalam 5

kelompok
Group

Environment

O2 Effect

Aerobic

Anaerobic

Obligate
Aerobe

Growth

No growth

Microaerophile

Growth
if
level not too No growth
high

Required but at levels

below 0.2 atm

Obligate
Anaerobe

No growth

Growth

Toxic

Required
(utilized
aerobic respiration)

for

Facultative
Anaerobe
(Facultative
Aerobe)

Growth

Growth

Not required for growth

but
utilized
when
available

Aerotolerant
Anaerobe

Growth

Growth

Not required
utilized

and

not

BAKTERI ANAEROB GRAM POSITIP

BERSPORA
[CLOSTRIDIUM spp]
Genus ini meliputi kuman berbentuk batang pleimorf, spora lebih
besar dari badan kuman, anaerob obligat
Gram-positive rods capable of forming endospores was placed in
the

genus

Clostridium.

This

genus

properties:
(1) presence of endospores,
(2) strict anaerobic metabolism,
(3) inability to reduce sulfate to sulfite, and
(4) Gram-positive cell wall structure

was

defined

by

four

 The organisms are ubiquitous in soil, water, and sewage and are part of the
normal microbial flora in the gastrointestinal tracts of animals and humans.
Most clostridia are harmless saprophytes
The majority of infections seen today are skin and soft-tissue infections, food
poisoning, and antibiotic-associated diarrhea and colitis.
The remarkable capacity of clostridia to cause diseases is attributed to their
(1)ability to survive adverse environmental conditions through spore formation;
(2)rapid growth in a nutritionally enriched, oxygen-deprived environment
(3)production of numerous histolytic toxins, enterotoxins, and neurotoxins.

Species

Human Disease

C. difficile

Antibiotic-associated diarrhea,
colitis
Soft-tissue infections (e.g., cellulitis, suppurative Common
myositis, myonecrosis, gas gangrene), food poisoning,
enteritis necroticans, septicemia
Gas gangrene, septicemia
Uncommo
n
Botulism
Uncommo
n
Tetanus
Uncommo
n
Opportunistic infections
Uncommo
n
Botulism
Rare
Botulism
Rare
Opportunistic infections
Rare

C. perfringens

C. septicum
C. botulinum
C. tetani
C. tertium
C. baratii
C. butyricum
C.
clostridioforme
C. histolyticum
C. innocuum
C. novyi
C. sordellii
C. sporogenes

Gas gangrene
Opportunistic infections
Gas gangrene
Gas gangrene
Opportunistic infections

Frequenc
y
pseudomembranous Common

Rare
Rare
Rare
Rare
Rare

Clostridium tetani

Differential diagnosa
Kuman berbentuk panjang langsing, agak bengkok, Gram +/P, ukuran
4,8 x 0,5, tunggal atau berbentuk rantai
Spora bulat, terminal [seperti pemukul genderang], tidak berkapsul dan
motil
Suhu pertumbuhan optimum 370C dan pH 7,4
Tumbuh baik pada media agar darah dengan zona hemolisa alfa
kemudian menjadi beta akibat pembuatan hemolisin [tetanolisin]

Toksin, membentuk 3 jenis toksin :

Hemolisin [tetanolisin]
Termolabil, tidak tahan terhadap oksigen, bekerja aktif pada SDM dari
sebagian besar binatang[ kelinci, kuda dll]
Peran belum jelas mungkin bekerja sebagai lekosidin

Neurotoksin [tetanospasmin]

Tahan terhadap oksigen dan diinaktifkan pada 66 0C selama 5 mnt. dapat

dibuat toksoid dengan penambahan formaldehid konsentrasi rendah
Bekerja seperti striknin dengan menghambat sintesis dan pelepasan
asetilkolin sehingga menganggu transmisi neuromuskuler. Zat ini akan
terikat kuat pada ganglion otak menyebabkan penghambatan neuron
spinal pasca sinaps dengan mencegah pelepasan perantara penghambat
[inhibitory mediator] kejang otot menyeluruh, reflek yang berlebihan
dan serangan kejang berulang.

Neurotoksin non-spasmogenik yang bekerja di saraf perifer

Gram stain of Clostridium tetani. Note the terminal spores.

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Clostridium tetani

Patogenesis

Kuman tidak bersifat invasif, akan tetap diluka, dapat

menjadi bentuk vegetatif bila kondisi
mendukung/anaerob, misalnya adanya:

Jaringan nekrotik

Garam kalsium

Kuman piogenik
Beberapa jenis tetanus :
1. Tetanus neonatorum : akibat pencemaran luka
pemotongan tali pusat bayi, angka kematian tinggi
2. Tetanus pasca keguguran dan masa nifas; akibat Infeksi
saluran kelamin oleh alat bantu persalinan dan pembalut
3. Splanchnic tetanus: terjadinya kekejangan otot otot
menelan dan pernafasan
4. Cephalic tetanus: luka di daerah kepala akan terjadi
kontraksi otot otot muka unilateral dan bilateral

Facial spasm and risus sardonicus in a patient with tetanus.

(From Cohen J, Powderly WG: Infectious diseases, ed 2, St Louis, 2004,
Mosby).
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A child with tetanus and opisthotonos resulting from persistent spasms of

the back muscles. (From Emond RT, Rowland HAK, Welsby P: Colour atlas
of infectious diseases, ed 3, London, 1995, Wolfe.)
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Clostridium tetani

Diagnosa laboratorium
Pemeriksaan mikroskopis: sampel diambil dari luka, diwarnai Gram
+/P seperti pemukul genderang
Biakan: potongan jaringan nekrotik dibiakkan pada kaldu daging/agar
darah
Percobaan binatang : untuk melihat toksigenitas kuman, digunakan
mencit yang disuntikan 0,2 ml suspensi kuman pada pangkal ekor,
kemudian dilihat mulai terjadinya kekejangan

Pencegahan & pengobatan

Pembersihan luka
Imunisasi aktif dengan toksoid
Imunisasi pasif dengan ATS
Pemberian antibiotik :Penicillin

PENCEGAHAN TETANUS PADA LUKA

Keadaan luka

Keadaan kekebalan
Kebal

Kebal sebagian Tidak kebal

Bersih [kecelakaan
rumah dlm 6 jam

1 x toksoid

1 x toksoid

3 x toksoid

Tercemar [tanah, ada

jaringan nekrotik]

1 x toksoid

1 x toksoid
ATS, Antibiotik

3 x toksoid
ATS, Antibiotik

Terinfeksi

1 x toksoid
Antibiotik

1 x toksoid
Ats, Antibiotik

3 x toksoid
ATS, Antibiotik

Catatan :
Kebal :Artinya telah menerima dosis lengkap berupa 3 x penyuntikan toksoid
Kebal sebagian :artinya telah menerima 2 x suntikan toksoid
Tidak kebal : artinya belum menerima toksoid atau keadaan kekebalan tidak
diketahui

Clostridium perfringens

Dahulu dikenal C. welchii, merupakan salah satu penyebab ganggren gas,

keracunan makanan oleh enterotoksin yang termolabil atau enteritis nekrotik

C. perfringens can be associated with simple colonization or can cause lifethreatening disease. C. perfringens is a large (0.6 to 2.4 1.3 to 19.0 m),
rectangular, Gram-positive rod , with spores rarely observed either in vivo or
after in vitro cultivation.

This organism is one of the few nonmotile clostridia, but rapidly spreading
growth on laboratory media (resembling the growth of motile organisms) is
characteristic (GAMBAR 2). The organism grows rapidly in tissues and in
culture, is hemolytic, and is metabolically active, features that make possible
its identification in the laboratory. The production of one or more major lethal
toxins by C. perfringens (alpha [], beta [], epsilon [], and iota [] toxins) is
used to subdivide isolates into five types (A through E; TABEL). Type A C.
perfringens causes most of the human infections in the United States.

Gram stain of Clostridium perfringens.

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GAMBAR 2. Growth of Clostridium perfringens on sheep blood agar. Note

the flat, spreading colonies and the hemolytic activity of the organism. A
presumptive identification of C. perfringens can be made by detection of
a zone of complete hemolysis (caused by the ;-toxin) and a wider zone
of partial hemolysis (caused by the ;-toxin), combined with the
characteristic microscopic morphology.
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Type

A
B
C
D
E

Lethal Toxins
Alph Beta Epsil
a
on
Iota
+
+
+
+
+
+
+
+
+
+

Distribution of Lethal Toxins in Clostridium perfringens

Types A to E

Growth of Clostridum perfringens on egg-yolk agar. The ;-toxin

(lecithinase) hydrolyzes phospholipids in serum and egg yolk,
producing an opaque precipitate (right). This precipitate is not
observed when the organism is grown in the presence of
antibodies against the toxin (left). This reaction (Nagler's reaction)
is characteristic of C. perfringens.

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Virulence
Factors
-Toxin

-Toxin
-Toxin
-Toxin

-Toxin
-Toxin
-Toxin
-Toxin
-Toxin
-Toxin
Enterotoxin

Biologic Activity
Lethal toxin; phospholipase C (lecithinase); increases
vascular
permeability; hemolysin; produces necrotizing activity, as
seen in
myonecrosis
Lethal toxin; necrotizing activity
Lethal toxin; permease
Lethal binary toxin; necrotizing activity; adenosine
diphosphate (ADP)
ribosylating
Hemolysin
Heat- and oxygen-labile hemolysin; cytolytic
Collagenase; gelatinase; necrotizing activity
Protease
Hyaluronidase
Deoxyribonuclease; hemolysin; necrotizing activity
Alters membrane permeability in ileum (cytotoxic,
enterotoxic);
superantigen

Pada ganggren gas, karbohidrat akan dihancurkan dengan

pembentukan gas karena ada septikemia, terjadi hemolisis
intravaskuler.

Pada keracunan makanan, enterotoksin merangsang enzim

adenylate cyclase pada dinding usus bertambahnya konsentrasi
cAMP hipersekresi air dan Cl dalam usus menghambat
reabsorpsi Na Diare [selama 1-3 hari]

Diagnosa laboratorium
Spesimen dari luka, pus, jaringan atau makanan
Mikroskopis: Gram +/P, batang tanpa spora
Biakan: di tanam pada agar tioglikolat, agar darah secara an aerob

Pengobatan
Pembersihan luka secara bedah pada jaringan nekrotik
Antibiotik : Penisilin
Hyperbaric oxygen

Clostridial cellulitis. Clostridia can be introduced into tissue during surgery

or by a traumatic injury. This patient suffered a compound fracture of the
tibia. Five days after the injury, the skin became discolored and bullae and
necrosis developed. A serosanguineous exudate and subcutaneous gas
were present, but there was no evidence of muscle necrosis. The patient
had an uneventful recovery. (From Lambert H, Farrar W, editors: Infectious
diseases illustrated, London, 1982,
DownloadedGower.)
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Clostridium botulinum

Terdapat secara luas di alam, kadang terdapat pada feses binatang

C. botulinum, the etiologic agent of botulism, is a heterogeneous
group of large (0.6 to 1.4 3.0 to 20.2 m), fastidious, sporeforming, anaerobic rods
Berdasarkan sifaT biokimia ada 2 jenis : Proteolitik [jenis A, B dan F],
jenis sakarolitik dan tidak proteolitik [Jenis C, D, dan E ] semua jenis
kelompok ini menghasilkan sulfida
Dosis letal untuk manusia : 1 mikrogram
Tidak menyebabkan infeksi luka, tetapi menyebabkan keracunan
makanan, akibat toksin yang termakan. Umumnya makanan yang
tercemar adalah makanan yang berbumbu, diasap, kalengan tanpa
dimasak terlebih dahulu.
Kerja toksin akan memblokir pembentukan/pelepasan acetyl cholin
pada hubungan saraf otot sehingga terjadi kelumpuhan otot.
Gejala 18-96 jam makan toksin dengan keluhan penglihatan, hal ini
terjadi karena tidak ada koordinasi. Sulit menelan, sulit bicara.
Kematian terjadi karena paralisis otot pernafasan atau kelumpuhan
jantung [cardiac arrest]
Other species of clostridia produce botulinum toxins, including C.
butyricum (type E toxin), C. baratii (type F toxin), and Clostridium
argentinense (type G toxin). Human disease has only rarely been
associated with C. butyricum and C. baratii, and not definitively
demonstrated with C. argentinense.

Group
I
II
III

Neurotoxin
Type
A, B, F
B, E, F
C, D

IV

Phenotyptic Properties
Proteolytic, saccharolytic
Nonproteolytic, saccharolytic
Weakly
proteolytic,
saccharolytic
Weakly
proteolytic,
asaccharolytic

Clostridium botulinum
Classification and Toxin
Production

Clostridium botulinum
Diagnosa laboratorium : mendeteksi toksin yang
ada didalam serum penderita atau dari sisa
makanan. Pendeteksian dilakukan dengan reaksi
Netralisasi antigen-antibodi secara agglutinasi
SDM yang dilapisis antiserum. Percobaan mencit
yang disuntikkan bahan tersangka.
Pengobatan:Pemberian antitoksin polivalen [tipe
A,B dan C] secara IV dan secara simptomatik
terutama untuk pernafasan
Pencegahan: Makanan yang diawetkan harus
dimasak dahulu secara baik, makanan yang perlu
diperhatikan : Kacang-kacangan, jagung, ikan
asap atau ikan segar dalam plastik

Clostridium botulinum

CLOSTRIDIUM DIFFICILE
Until the mid-1970s the clinical importance of C.

difficile was not appreciated. This organism was

infrequently isolated in fecal cultures and its role in
human disease was unknown. Systematic studies
now clearly show, however, that toxin-producing C.
difficile is responsible for antibiotic-associated
gastrointestinal diseases , ranging from a relatively
benign,

self-limited

diarrhea

to

severe,

life-

threatening pseudomembranous colitis (GAMBAR).

Antibiotic-associated colitis: gross section of the lumen of the colon. Note

the white plaques of fibrin, mucus, and inflammatory cells overlying the
normal red intestinal mucosa.
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Antibiotic-associated colitis caused by Clostridium difficile. A histologic

section of colon shows an intense inflammatory response, with the
characteristic "plaque" (black arrow) overlying the intact intestinal
mucosa (white arrow). (Hematoxylin and eosin stain.)
(From Lambert HP, Farrar WE, editors: Infectious diseases illustrated,
London, 1982, Gower.)
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Virulence Factor
Enterotoxin (toxin A)

Cytotoxin (toxin B)

Adhesin factor
Hyaluronidase
Spore formation

Biologic Activity
Produces chemotaxis; induces
cytokine
production
with
hypersecretion
of
fluid;
produces hemorrhagic necrosis
Induces depolymerization of
actin with loss of cellular
cytoskeleton
Mediates binding to human
colonic cells
Produces hydrolytic activity
Permits organism's survival for
months
in
hospital
environment

. Virulence Factors Associated with Clostridium

difficile

ANAEROB, GRAM POSITIP,

TIDAK MEMBENTUK SPORA
The anaerobic gram-positive cocci and non- spore-forming rods are a
heterogeneous group of bacteria that characteristically colonize the
skin and mucosal surfaces.
These organisms are opportunistic pathogens, typically responsible
for endogenous infections and usually recovered in mixtures of
aerobic and anaerobic bacteria.
Most of these anaerobes have fastidious nutritional requirements and
grow slowly on laboratory media. Thus the isolation and identification
of individual strains are difficult and often time consuming.

Organism
Anaerobic Cocci
Anaerococcus
Finegoldia
Micromonas
Peptostreptococcus
Schleiferella
Anaerobic Rods
Actinomyces
Bifidobacterium
Eubacterium
Lactobacillus

Mobiluncus
Propionibacterium
propionicum

Historical Derivation
an, without; aer, air; coccus, berry or coccus
(anaerobic coccus)
Named after the American microbiologist S. Finegold
micro, tiny; monas, cell (tiny cell) pepto, cook or
digest (the digesting streptococcus)
Named after the German microbiologist K.H. Schleifer
aktinos, ray; mykes, fungus (ray fungus referring to
the radial arrangement of filaments in granules)
bifidus, cleft; bakterion, small rod (a small clefted or
bifurcated rod)
eu, good or beneficial (a beneficial rod; that is, a rod
normally present)
acto, milk (milk bacillus; organism originally recovered
in milk; also, lactic acid is the primary metabolic
product of fermentation)
mobilis, capable of movement or being active; uncus,
hook (motile, curved rod)
propionic acid (propionic acid is the primary metabolic
product of fermentation)

Important Anaerobic Gram-Positive

Bacteria

ACTINOMYCES

Actinomyces organisms are facultatively anaerobic or strictly anaerobic, gram-positive

rods. They are not acid-fast (in contrast to the morphologically similar Nocardia species),

They grow slowly in culture, and they tend to produce chronic, slowly developing
infections. They typically develop delicate filamentous forms or hyphae (resembling fungi)
in clinical specimens or when isolated in culture (GAMBAR.

However, these organisms are true bacteria in that they lack mitochondria and a nuclear
membrane, reproduce by fission, and are inhibited by penicillin but not antifungal
antibiotics. Numerous species have been described; Actinomyces israelii, Actinomyces
meyeri, Actinomyces naeslundii, Actinomyces odontolyticus, and Actinomyces viscosus
are responsible for most human infections.

Only A. meyeri is a strict anaerobe. The other species grow best in anaerobic conditions
but can grow aerobically.

Macroscopic colony (left) and Gram stain (right) of Actinomyces.

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PATHOGENESIS AND IMMUNITY

Actinomyces

organisms

colonize

the

upper

respiratory,

gastrointestinal, and female genital tracts. These bacteria are not

normally present on the skin surface. The organisms have a low
virulence potential and cause disease only when the normal mucosal
barriers are disrupted by trauma, surgery, or infection.

Disease caused by actinomyces is termed actinomycosis (in keeping

with the original idea that these organisms were fungi or "mycoses").
Actinomycosis is characterized by the development of chronic
granulomatous lesions that become suppurative and form abscesses
connected

by

sinus

tracts.

Macroscopic

colonies

of

organisms

resembling grains of sand can frequently be seen in the abscesses and

sinus tracts. These colonies, called sulfur granules because they
appear yellow or orange, are masses of filamentous organisms bound
together by calcium phosphate (GAMBAR).

Sulfur granule collected from the sinus tract in a patient with

actinomycosis. Delicate filamentous rods (arrow) are seen at the periphery
of the crushed granule.
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EPIDEMIOLOGY

Actinomycosis is an endogenous infection

Patients with thoracic infections generally

person-to-person

have a history of aspiration, with the disease

spread or disease originating from an

becoming established in the lungs and then

with

no

evidence

of

external source, such as soil or water.

Disease is classified according to the organ

systems involved. Cervicofacial infections
are seen in patients who have poor oral
hygiene or have undergone an invasive
dental procedure or oral trauma. In these
patients, the actinomyces that are present
in the mouth invade into the diseased
tissue and initiate the infectious process.

spreading to adjoining tissues. Abdominal

infections most commonly occur in patients
who have undergone gastrointestinal surgery
or have suffered trauma to the bowel. Pelvic
infection can be a secondary manifestation of
abdominal actinomycosis or may be a primary
infection in a woman with an intrauterine
device (Central nervous system infections
usually represent hematogenous spread from
another infected tissue, such as the lungs.

TREATMENT, PREVENTION, AND CONTROL

Treatment

for

actinomycosis

involves

the

combination

surgical

dbridement

of

of
the

The

clinical

generally

response

good

even

is
in

involved tissues and the prolonged

patients who have suffered

administration of antibiotics.

extensive tissue destruction.

Actinomyces

are

uniformly

susceptible

to

penicillin

(considered
choice),
clindamycin.

the

antibiotic

erythromycin,
Most

species

of

Maintenance
hygiene

and

of

good

the

use

oral
of

and

appropriate

are

prophylaxis when the mouth

resistant to metronidazole, and the

tetracyclines have variable activity.

or

gastrointestinal

antibiotic

tract

is

penetrated can lower the risk

of these infections.

Propionibacterium

Propionibacteria are small gram-positive rods often arranged in

short chains or clumps

They are commonly found on the skin (in contrast with the
actinomyces), conjunctiva, external ear, and in the oropharynx
and female genital tract. The organisms are anaerobic or
aerotolerant,

nonmotile,

catalase

positive,

and

capable

of

fermenting carbohydrates. The two most commonly isolated

species

are

propionicus.

Propionibacterium

acnes

and

Propionibacterium

P. acnes is responsible for two types of infections:

(1) acne (as the name implies) in teenagers and young adults and
(2) opportunistic infections in patients with prosthetic devices (e.g., artificial heart
valves or joints) or intravascular lines (e.g., catheters, cerebrospinal fluid shunts).

Propionibacteria are also commonly isolated in blood cultures, but this finding
usually represents contamination with bacteria on the skin at the phlebotomy site.

Acne is unrelated to the effectiveness of skin cleansing because the lesion develops
within the sebaceous follicles. For this reason, acne is managed primarily through
the topical application of benzoyl peroxide and antibiotics.

Antibiotics such as erythromycin and clindamycin have proved effective for

treatment.

Mobiluncus

Members of the genus Mobiluncus are

obligate

anaerobic,

gram-variable

or

growing

gram-negative, curved rods with tapered

because they

even

on

with rabbit or horse serum.

stained specimens
they are classified as gram-positive rods

slowly

enriched media supplemented

ends. Despite their appearance in Gram-

The organisms are fastidious,

Two

species,

curtisii

and

Mobiluncus
Mobiluncus

(1) have a gram-positive cell wall,

mulieris, have been identified

(2) lack endotoxin, and

in

(3) Are susceptible to vancomycin, clindamycin,

colonize the genital tract in

erythromycin, and ampicillin but resistant to

low numbers but are abundant

colistin.

in

humans.

women

The

with

organisms

bacterial

vaginosis (vaginitis).

Lactobacillus

Lactobacillus

species

are

facultatively anaerobic or strictly

anaerobic rods.

They are found as part of the

normal

flora

stomach,
genitourinary

of

the

mouth,

intestines,

and

tract.

The

organisms are most commonly

isolated in urine specimens and
blood

cultures.

Because

lactobacilli are the most common

organism in the urethra

The

reason

lactobacilli

rarely

cause infections of the urinary

tract is their inability to grow in
urine. Invasion into blood occurs
in one of the following three
settings: (1) transient bacteremia
from a genitourinary source (e.g.,
after childbirth or a gynecologic
procedure), (2) endocarditis, and
(3) opportunistic septicemia in an
immunocompromised patient.

Lactobacillus
Treatment

of

endocarditis

and

opportunistic infections is difficult because

lactobacilli are resistant to vancomycin (an
antibiotic commonly active against grampositive bacteria) and are inhibited but not
killed by other antibiotics. A combination
of penicillin with an aminoglycoside is
required for bactericidal activity.

Bifidobacterium and Eubacterium

Bifidobacterium and Eubacterium species
are commonly found in the oropharynx,
large

intestine,

bacteria

can

be

and

vagina.

isolated

in

These
clinical

specimens but have a very low virulence

potential and usually represent clinically
insignificant contaminants.

Anaerobic Gram-Negative
Bacteria

The most important gram-negative anaerobes

that colonize the human upper respiratory,
gastrointestinal, and genitourinary tracts are
the

rods

in

the

genera

Bacteroides,

Fusobacterium, Porphyromonas, and Prevotella

and the cocci in the genus Veillonella
Among these pathogens, the most important is
Bacteroides

fragilis,

the

endogenous anaerobic pathogen.

prototypical

Table . Virulence
Negative Rods
Virulence Factor
Adhesins
Capsule

Factors

in

Anaerobic

Gram-

Bacteria
Bacteroides
fragilis,
Prevotella
melaninogenica
B. fragilis, Porphyromonas gingivalis
P. gingivalis
Fusobacterium nucleatum

Fimbriae
Hemagglutinin
Lectin
Resistant to Oxygen Toxicity
Superoxide dismutase
Many species
Catalase
Many species
Antiphagocytic
Capsule
B. fragilis, P. melaninogenica
Immunoglobulin (Ig)A, IgM, IgG Porphyromonas spp.,
proteases
Prevotella spp.
Lipopolysaccharide
Fusobacterium sp
Succinic acid
Many species

Virulence Factor
Tissue Destruction
Phospholipase C
Hemolysins
Proteases
Collagenase
Fibrinolysin
Neuraminidase
Heparinase
Chondroitin sulfatase
Glucuronidases
N-Acetylglucosaminidase
Volatile fatty acids
Toxin
Enterotoxigenic toxin

Bacteria
Fusobacterium necrophorum
Many species
Many species
Many species
Many species
Many species
Many species
Many species
Many species
Many species
Many species
B. fragilis

Modified from Duerden B: Clin Infect Dis 18(suppl 4):S253-S259, 1994; and Lorber B: Bacteroides,
Prevotella, Porphyromonas, and Fusobacterium species. In Mandell, Douglas and Bennett's principles
and practice of infectious diseases, ed 6, New York, 2005, Churchill Livingstone.